Apidologie (2015) 46:10–22 Review article

* The Author(s), 2014.

This article is published with open access at Springerlink.com
DOI: 10.1007/s13592-014-0298-x

Application of continuous monitoring

of honeybee colonies

W. G. MEIKLE1 , N. HOLST2
1
USDA-ARS, 2000 E. Allen Rd, Tucson, AZ 85719, USA
2
Department of Agroecology, Aarhus University, Forsøgsvej 1, 4200 Slagelse, Denmark

Received 7 January 2014 – Revised 24 April 2014 – Accepted 3 June 2014

Abstract – Monitoring physical variables associated with honeybee colonies, including weight, temperature,
humidity, respiratory gases, vibration, sound, and forager traffic, in a continuous manner is becoming feasible
for most researchers as the cost and size of electronic sensors decrease while their precision and capacity
increase. Researchers have taken different approaches to collecting and analyzing the resulting datasets, with a
view toward extracting information on colony behavior and phenology. The objective of this review is to
examine critically the different kinds of data and data analyses, providing researchers with better-informed
options for obtaining information on colony phenology in the field without disturbing the hive, and for
combining information from different kinds of sensors to obtain a more complete picture of colony status.
Wireless sensor networks and powering sensors are briefly discussed.

continuous hive weight / colony temperature / colony humidity / forager traffic / hive vibration

1. INTRODUCTION remotely, and with little manpower. Once sensors

Interest in monitoring honeybee colonies on a
continuous basis, defined here as data gathered from
the colony (as opposed to individual bees) hourly or
more often for periods exceeding 2 days, is not new.
Gates (1914), for example, reported hourly temper-
ature data over several days collected from a
beehive in 1907. However, sensor technology has
changed a great deal, and its application to both bee
research and general beekeeping is increasing.
Smaller, cheaper, and more accurate sensors, along
with easier connections to computers and the
Internet (Faludi 2010), have made it possible for
bee researchers and beekeepers to monitor many
physical aspects of bee colonies continuously,
Corresponding author: W.G. Meikle,
william.meikle@ars.usda.gov
Manuscript editor: Peter Rosenkranz
have been installed, hives can be monitored without
disturbance, including during periods when invasive
hive inspections are contraindicated, such as during
winter or times of colony stress.
Honeybee colonies present particular advan-
tages as subjects for continuous monitoring,
especially when kept in hives where all parts of
the colony are easily separated, measured, and, if
need be, modified. Honeybee colonies are active,
in one way or another, all day every day and have
been considered as “superorganisms” (Southwick
and Mugaas 1971; Moritz and Southwick 1992;
Schmolz et al. 1994) with individual bees playing
roles analogous to cells in a multicellular organism.
Bees behave collectively in ways that isolated or
solitary bees do not, such as by regulating
temperature and humidity (Human et al. 2006),
and by protecting the colony against diseases and
pests (Evans and Spivak 2010). How effectively a
given colony performs these collective, complex
behaviors can be used as indicators of colony
 Application of continuous monitoring of honeybee colonies
genetics, phenology, and health. The capacity of a
colony to regulate colony temperature, for exam-
ple, has been found to be a function of the bee
subspecies (W-Worswick 1987), the within-colony
genetic diversity (Jones et al. 2004), and pheno-
logical status (Stalidzans and Berzonis 2013).
Healthy honeybee colonies maintain, in many
respects, a stable environment within at least part of
the hive, but that environment is still subject to
changing biotic and abiotic factors. A local nectar
flow can rapidly boost the colony’s food reserves
(Gary 1992), or a sudden exposure to pesticide in a
field nearby may suddenly reduce the number of
foragers (Rortais et al. 2005). A sudden period of
unusually cold weather may reduce forager activity
and brood production (Hoopingarner and Waller
1992). Internal events, such as the death of the queen
or reproductive swarming, may cause abrupt chang-
es to colony dynamics (Gary 1992). By monitoring
colonies continuously, a researcher conducting field
experiments can account for such factors when
evaluating treatment effects. Studies that addressed
bee health and behavior only outside the hive, such
as foraging activity measured at flowers (e.g., Sabara
and Winston 2003) or at feeding dishes (e.g., Colin
et al. 2004), were not considered here.
Here, we present early and recent studies
employing continuous monitoring of physical
parameters of honeybee colonies and discuss
method application and data interpretation.
Continuous monitoring provides longitudinal data
that allows correlation of hive events, such as
changes in forager activity, with changes in hive
health, phenology, and queen status, and with
external factors, such as weather, nectar flow, or
pesticide exposure, and it provides an important
perspective to studies on the interactions between
colony health and the environment.
2. LITERATURE REVIEW
The objectives, methods, location (field or
laboratory), and duration of studies involving
continuous monitoring of bee colonies have
varied among researchers (Table I). Sensors have
been grouped here into four main types: (1)
weight; (2) temperature, humidity, and gas; (3)
sound and vibration; and (4) forager traffic. The
11
variables examined in these studies can be
considered either state variables (weight, temper-
ature, humidity, gases) or rate variables (forager
traffic), which offer different options for statistical
analysis and biological inference. Among the state
variables, colony weight at any particular moment
is merely a physical characteristic with little
information on colony status per se, but first-
and second-order changes in weight over time are
informative. Temperature, humidity, and respira-
tory gas concentrations are somewhat different
since these variables are directly related to the
metabolism of the bee colony (Kronenberg and
Heller 1982; Van Nerum and Buelens 1997).
There are often temperature and gas gradients
within a hive, so data depend on the location and
precision of the sensor. Vibration and sound are
difficult to categorize as state or rate as they can be
considered in both time and frequency domains.
Bees use vibration and sound to communicate, but
they also produce vibrations and sounds they
likely do not use (Atauri Mezquida and Llorente
Martínez 2009) and may not even detect, although
those may provide information about the hive. A
final section discusses the application of wireless
networks to continuously monitored systems.
2.1. Colony weight
Placing a honeybee hive on a scale to weigh it
disturbs the hive very little—and if the hive is kept
on the scale, then weighing it does not disturb the
bees at all. Occasional weighing (weekly or even
daily) is usually done to determine when to
harvest honey or to estimate hive food reserves
(e.g., McLellan 1977; Szabo and Lefkovitch
1991; Harbo 1993). Continuous weighing with a
sufficiently precise scale can provide that infor-
mation as well as data on shorter-term changes in
the hive. Weight data is easy to define and
analyze: a colony has a single weight value at a
given point in time, and scales are widely
available and easily installed. Most load cells
control for temperature variability, at least over a
given range of values, but some weather factors,
such as precipitation and wind, can affect the data.
In early applications of continuous monitoring,
Gates (1914) and Hambleton (1925) placed 10-
Table I. Summary of research using continuous monitoring methods. 12

Parameter Method Location No. of replicate Duration of hourly Reference

colonies datasets

Weight and Mechanical balance and in-hive Field 1 1 year: hourly during Gates (1914)
temperature mercury thermometers the day
Weight Mechanical balance Field 2 12 days Hambleton (1925)
Weight Electronic balance Field 1 1 month Buchmann and Thoenes
(1990)
Weight Electronic balance Field 2–4 16 months Meikle et al. (2006, 2008)
Temperature Electric thermocouples Insulated room 1 5 months; hourly during Phillips and Demuth (1914)
the day
Temperature, [O2] and Electric thermocouples, metabolic Field 1 3 days Milner (1921)
[CO2] chamber with extracted air
passed through external
detectors
Temperature Extracted air passed over Field 1 Approx. 5 days Southwick and Mugaas
thermometer (1971)
Temperature, [CO2], In-hive mercury thermometers, Not stated 1 52 h Seeley (1974)
no. of fanning bees extracted air passed through
external detectors
W.G. Meikle and N. Holst

Temperature, [O2], no. Metabolic chamber with Laboratory Not stated Series of 24–48-h Kronenberg and Heller
of fanning bees extracted air passed through experiments over (1982)
external detectors 14 months
Temperature, [O2] and Extracted air passed through Field 10 Not specified; some periods Southwick and Moritz
[CO2] detectors of at least 12 h (1987)
Temperature, [O2] and In-hive temperature sensors; Field and Not stated 18 weeks Van Nerum and Buelens
[CO2], humidity extracted air passed through laboratory (1997)
gas detectors
Temperature In-hive sensors Field 8 2 weeks Jones et al. (2004)
Temperature, humidity In-hive sensors Field 3 (incl. empty 4 days Human et al. (2006)
control)
Temperature In-hive sensors Field 14 1 year Stalidzans and Berzonis
(2013)
Vibration In-hive sensors Field 2 Approx. 8 months Bencsik et al. (2011)
 Application of continuous monitoring of honeybee colonies 13

Danka and Beaman (2007)

frame Langstroth hives on mechanical balances in

Burrill and Dietz (1981)

sheltered locations. About every 3 weeks from

Schneider et al. (2012)

Atauri Mezquida and
October 1907 to September 1908, Gates (1914)
Ferrari et al. (2008)

Llorente Martínez
Reference

recorded “by means of assistance” weight and

temperature data hourly for 2 to 3 days but presented
data on daily weight change only for the winter,
(2009)

including changes caused by rainfall. Hambleton

(1925) established a team of three people, in 8-h
shifts, to manually record the hourly hive weight
changes for several days in September 1922, and for
13 consecutive days in May 1923. He noted the
Duration of hourly

strong daily pattern of weight loss, attributed to

water loss through nectar drying and respiration
10 trials of up to
Approx. 1 year

during the night and to departing foragers early in

datasets

48 h each

the morning, followed by weight gain due to

foraging from mid-morning until late evening.
23 days
9 days
270 h

Hambleton (1925) correlated ambient weather

variables, including temperature, relative humidity,
and hours of sunshine, with hive weight changes,
No. of replicate

under the assumption that those factors affect nectar

production which in turn affects colony weight.
colonies

Use of manual labor by Gates (1914) and

Hambleton (1925) to record weight data was
10

40
3

dictated by the technology available at the time.

Buchmann and Thoenes (1990) introduced the use
of continuous weight monitoring using a precision
Location

scale linked to a computer to record data every

15 min for 1 month (August 1988) for a single hive
Field

Field

Field
Field
Field

and used the data to link hive abandonment to

tracheal mite (Acarapis woodi Rennie) infestation.
Meikle et al. (2006, 2008) deployed a similar
system: hives placed on precision electronic scales
linked to data loggers. Similar to previous workers,
RFID tags and entrance

data loggers were set to record weight hourly or

Hive entrance sensors
Hive entrance sensors
Method

more often, and weight was recorded for up to four

hives over 16 months. Meikle et al. (2006, 2008)
In-hive sensors

In-hive sensors

calculated the 25-h running average, subtracted

sensors

those values from the hourly raw data to produce

“detrended” residuals and then fit sine functions to
these residuals. They showed the effects of
swarming on daily detrended data, and also that
Acoustics, temperature,

Acoustics, temperature

an empty wooden hive has a detectable daily

Table I (continued)

relative humidity

weight change pattern, probably due to moisture

content changes in the wood.
Forager traffic
Forager traffic
Forager traffic

Continuous weighing has been shown to pro-

Parameter

vide information on weather effects (Gates 1914;

Hambleton 1925; Meikle et al. 2006), colony
growth and consumption (Meikle et al. 2008),
14 W.G. Meikle and N. Holst
swarming (Buchmann and Thoenes 1990; Meikle
et al. 2006), hive abandonment (Thoenes and
Buchmann 1992), the impact of pesticides on bee
colonies, changes in nectar and pollen availability,
and differences among honeybee races (Buchmann
and Thoenes 1990). Continuously weighing hives
provides data on colony-level weight changes but
not with respect to colony component (adults,
brood, and food reserves), and while detrended
data shows forager activity, this data is confounded
at least to some degree with water and pollen gain
and loss, which can be significant at certain times
of the year. For this reason, some measurement of
colony size and food reserves is useful for
interpreting weight data, although many studies
have not provided this. Meikle et al. (2008)
evaluated hives every 2 weeks over 6 months in
2005 and 3 months in 2006. By subtracting the
weights of the hive parts from the total hive weight,
and analyzing the photographs taken of each side
of each frame to determine the percentage of
capped brood and food reserves, they estimated the
masses of the adult bee and brood populations, the
food reserves, and the daily food demand. Changes
in the running average hive weight were correlated
with colony growth, and the amplitude of sine
waves fit to the detrended data was correlated with
daily colony food demand.
2.2. Colony temperature, humidity,
and gas concentrations
Like weight data, temperature, humidity, and
gas measurements provide single values per sensor
at a given point in time, but unlike weight data,
there is a range of values within a beehive,
depending on the number and placement of the
sensors. The position within the hive of a given
temperature or gas sensor will determine to what
extent it is affected by exterior ambient conditions,
and differences among various points within a hive
can be large. Temperature sensors, for example,
within the brood cluster, usually the warmest part
of the hive (Southwick 1992), will be affected less
by exterior conditions while those nearer the
exterior of the hive can be expected to be affected
more. Sensors at fixed points may change over
time with respect to their distance to the brood
cluster as bees move and change the size of the
cluster during the year and reduce or eliminate
them in the winter (Szabo 1989) when gradients in
temperature, for example, between exterior and
interior, and top and bottom, are more extreme.
Arrays of sensors embedded on the surface of hive
frames would solve this problem but may be
expensive to implement; Owens (1971) described
the use of thermocouple arrays in winter hives to
monitor cluster location and size. Researchers
without access to sensor arrays may need to
consider involving replicate hives in order to
control for variability in brood location. Sensor
placement can be standardized with respect to the
hive, such as on top of a middle frame in the brood
box, or with respect to the colony, such as the
center of the brood cluster. In the latter case,
sensors may be in a different place in each hive.
Gases (including water vapor) have been
sampled in beehives primarily using one of three
methods: (1) placing the colony in a metabolic
chamber and passing respiratory gases of known
composition through the colony while measuring
[O2] and [CO2] in the output (e.g., Milner 1921);
(2) removing air samples from the hive using
either pipettes (e.g., Van Nerum and Buelens
1997) or plastic tubing (e.g., Seeley 1974;
Southwick and Moritz 1987) and then measuring
gas in detectors outside the hive; or (3) placing
sensors within the hive (e.g., Human et al. 2006).
The first approach is ideal for colony-level studies
of gas exchange rather than studies of within-
colony gradients. The second approach allows
researchers to sample air at very precise locations
within the hive but will likely require hive
modification or some disturbance and may be
difficult to implement over large numbers of
hives. The third approach is convenient in that
commercially available gas sensors can fit easily
between or within frames and large numbers of
hives can be monitored simultaneously. However,
in the hive, bees tend to cover foreign objects with
propolis or wax which would interfere with air
movement across the sensor, so in-hive gas
sensors must be checked with some regularity to
ensure that they or their protective covers are
sufficiently clean. Care must also be taken when
placing sensors directly on brood, as the sensor
 Application of continuous monitoring of honeybee colonies
can interfere with comb maintenance and with bee
emergence (WGM pers. obs.). Data from tubes or
sensors at fixed sites may also be affected by
movement of the brood cluster.
Measurement of temperature and gas concentra-
tions has been conducted largely to understand
metabolic processes on the colony level. Milner
(1921), after making calculations from temperature
and O2 consumption, determined that the energy
produced by a colony of bees, even in conditions of
favorable temperatures and low disturbance,
exceeded that, when considered per unit weight,
of a “man when working at hard manual labor” and
expected that the energy production would be even
greater under less favorable conditions. Southwick
and Mugaas (1971) confirmed observations by
Gates (1914) that when ambient temperatures were
<5 °C, decreasing ambient temperature was associ-
ated with increasing colony core temperature
(mostly 20–32 °C). Seeley (1974) observed chang-
es in the number of fanning bees when [CO2], [O2],
and [N2] were manipulated; only changing [CO2]
influenced fanning behavior, presumably because
bees can detect changes in [CO2] but not the other
gases (Southwick and Moritz 1987). Van Nerum
and Buelens (1997) observed that bees actively
maintained low (15 %) O2 levels, which caused a
reversible hypoxia and reduced metabolic rate
among the bees which, they hypothesized, allowed
bees to combine water conservation, energy con-
servation, and longevity with the ability to increase
energy consumption on short notice if needed for
hive defense. By measuring bee fanning activity
while manipulating temperature, Jones et al. (2004)
found that temperature variation within a colony
was significantly lower in diverse patriline colonies
compared to the uniform patriline colonies, sug-
gesting that genetic diversity among workers may
promote greater temperature stability owing to
different genetic thresholds for action.
Several research groups have observed daily or
shorter-term temporal patterns in the data.
Kronenberg and Heller (1982) noted “periodic,
pulsatile increases in metabolic rate on the overall
daily rhythm” of the bee groups which they thought
might be linked to forager activity, and Southwick
and Moritz (1987) presented data on the change in
[O2] over 24 h, showing a clear diurnal cycle in the
15
frequency of respiration in addition to the higher-
frequency cycles of air movement. On an even finer
time scale, Southwick and Moritz (1987) observed
“peaks of tidal air movement” with a frequency of
about 22 s, and irregular periods of 4 to 37 min of
lower amplitude superimposed on the shorter
periods. To exploit daily patterns, Human et al.
(2006) applied cosinor analysis to the half-hour
averages in order to evaluate whether relative and
absolute humidity changes exhibited circadian
rhythmicity and whether that rhythmicity differed
among hives. Cosinor analysis involves fitting
sinusoidal waves to datasets using least sums-of-
squares and making statistical comparisons of wave
parameters, such as amplitude, phase, and mesor
(value about which an oscillation occurs), among
treatment groups (Nelson et al. 1979). Human et al.
(2006) found that bees can maintain absolute
humidity levels well above external levels, with
greater variability near the brood than near the
nectar and honey stores.
2.3. Hive vibration and sound
Vibration and sound are physically linked
phenomena measured at a point in space over
time, either on the surface of the hive or within the
hive volume. Vibration and sound data contain
rich spectra of overlapping wave forms and
require processing to distil biologically meaning-
ful information. Vibration of the hive substrate is
an important form of communication among bees
(Nieh and Tautz 2000; Sandeman et al. 1996;
Schneider et al. 1986), which have specialized
receptors in their legs for receiving low-frequency
signals, generally <300 Hz (Sandeman et al.
1996), and the signals are thought to involve
recruitment for foraging (Nieh and Tautz 2000).
However, bees in a hive produce many frequen-
cies of vibrations, from <10 to >1,000 Hz
(Bencsik et al. 2011), and how much of this
spectrum is used by bees is unknown. Several
methods have been used to gather vibration data.
Nieh and Tautz (2000) used laser vibrometry,
focusing a laser on the wall of a comb cell next to
a dancing bee and measuring the displacement.
The method provided accurate data for single bees
in a specific part of the comb, but is impractical
for use on entire bee colonies in the field.
16 W.G. Meikle and N. Holst
Vibration data for a beehive, when examined as
amplitude over time, tend to be “noisy” and contain
many peaks of different sizes (please see Figure 5 in
Nieh and Tautz 2000). One method to analyze this
kind of data is to apply a discrete Fourier transform,
which represents the data in the frequency domain
as the sum of a series of sinusoidal waves of
varying frequencies and amplitudes (Smith 2003)
and thereby obtain a frequency spectrum.
Amplitude data in both the time and frequency
domains can be represented on spectrograms and
those spectrograms examined for particular time
periods, such as waggle dancing (Nieh and Tautz
2000). Researchers have used two main approaches
to process vibration and sound data: (1) apply
principal components analysis (PCA) to the entire
range of resulting frequencies, and then evaluate the
components that capture most of the information,
and (2) focus on the amplitudes of a few frequency
bands identified beforehand.
Bencsik et al. (2011) embedded a vibration
sensor, an accelerometer, in the wall of each of
two hives, linking the sensor to a signal conditioner
and a computer with considerable disk space. They
collected data almost continuously from November
2008 to June 2009. Individual bees can be expected
to vibrate at different frequencies and without phase
coherence, so to resolve that problem, Bencsik et al.
(2011) maintained a frequency resolution of 20 Hz
and examined relatively short data samples of
510 s. They found that those samples produced
power spectra with a satisfactory signal to noise
ratio to determine frequency peaks, and they filtered
much of the data noise by first expressing data in
the frequency domain PCA and then selecting the
components with eigenvalues in the top 5 %.
Principal components are orthogonal to each other,
so each component represents a unique source of
variance. Bencsik et al. (2011) were thus able to
focus on the strongest patterns without losing
appreciable resolution; they used the PCA-filtered
data to examine vibration frequencies prior to bee
colony swarming.
Sound data is as rich as vibration data, but bee-
produced sounds are somewhat better understood.
Most beekeepers listen to their hives, and the
meaning of some sounds is well known (Schlegel
et al. 2012; Seeley 2010). Rangel and Seeley
(2008) took advantage of this by embedding a
microphone in each of five small (three-frame)
observation hives and, over the course of about a
month, used the microphones to detect a particular
kind of sound, “piping”. When piping was
detected at a certain threshold rate, in this case at
least three signals in 30 s, they switched on video
equipment to capture images that were later
analyzed for bee movement.
Long-term sound monitoring of entire hives has
seldom been reported, probably because the
resulting amount of data tends be very large and
unwieldy. Ferrari et al. (2008) monitored sound,
temperature and relative humidity for 270 hours
using within-hive sensors and observed 9
swarming events among three colonies; they
observed increases in sound intensity and drops in
temperature and humidity during the swarming.
Atauri Mezquida and Llorente Martínez (2009)
placed microphones and temperature sensors in up
to 10 hives between May 2008, and April 2009.
They recorded temperature within and outside the
hive, and for each 8-s sound sample, they
monitored the power (in W), the “rugosity” (an
index of noisiness), and the tone and intensity of
the five main sound frequency bands, as deter-
mined by a Fourier transform of the sound sample.
These analyses provided some qualitative data
concerning sound differences between healthy
hives and those infected with chalkbrood
(Ascosphaera apis), but the main goal of the paper
was to introduce methods to analyze continuous
sound data.
2.4. Forager traffic
Foraging is necessary for food gathering and is
generally conducted by older bees (Gary 1992;
Gould and Gould 1988). Forager traffic will thus
be affected by food availability, food demand, and
colony age structure (McLellan 1977), and sudden
changes in that traffic may indicate acute changes
on the colony level. Pham-Delègue et al. (2002)
described forager activity as an important variable
to monitor when evaluating the impact of pesti-
cides on honeybee colony health.
Forager activity is described in terms of the
number of bees entering and/or exiting the hive over
 Application of continuous monitoring of honeybee colonies
a given time period, and so data can be collected, if
need be, without the use of equipment more
sophisticated than an observer and a stopwatch.
Kolmes and Sam (1990) observed hives for several
minutes per day over 23 days to analyze bee activity
during hive establishment, and Corbett et al. (1993)
collected data visually on the activity of several
species of social bees with forager counts for several
hours per day over 1–4 days. The use of human
observation, while likely accurate, clearly limits,
due to fatigue, the amount of time that the hive can
be observed. Automating this has been a goal for
some time; Faberge (1943) proposed an electrical
counter as an improvement upon one originally
described by Lundie (1925), which depended on
bees tripping a balance arm, which produced
electrical impulses that drove a printer. Faberge
discussed the use of photovoltaic cells, but
dismissed them as too expensive at the time. Later,
Spangler (1969), Erickson et al. (1975), Liu et al.
(1990) and others developed such counters.
Several designs of hive entrance counters have
been used. Burrill and Dietz (1981) gathered
forager traffic data, using a photoelectric bee
counter (“Apicard”) placed at the hive entrance,
for 30-min periods (data were recorded every 15 s
and were pooled) over 23 consecutive days from a
single colony of bees and analyzed those data with
respect to ambient weather conditions. Danka and
Beaman (2007) used commercially available bee
counters of a design described by Struye et al.
(1994) to measure flight activity among 40
colonies of “Russian” and “Italian” bee races
during blueberry pollination. Part of the objective
of that study was to determine whether either race
conducted more foraging flights per hour than the
other race after controlling for weather and black
globe temperature (Corbett et al. 1993). Danka
and Beaman (2007) did observe a problem with
counters installed at the hive entrance: under
certain conditions, bees tend to cluster around
the sensor, causing spurious “entrance” and “exit”
readings, which can have large effects on the data,
so they conducted visual counts of forager flights
as well. Validating the hive entrance data using
other means, as Danka and Beaman (2007) did,
greatly increases ease of data interpretation.
17
Streit et al. (2003) introduced the use of radio-
frequency identification (RFID) tags to monitor
honeybee movement in and out of the hive. The
tags, which are small and weigh a few milligrams or
less, are attached to adult bees. A detector records
each time the tag crosses a threshold. To determine
whether a given bee is entering or leaving, two
detectors are used in one of two ways: (1) bees are
restricted to a single entrance hole and two detectors
arranged so that the order in which the bee crosses
the detector determines direction or (2) the entrance
is modified so that entering bees are obliged to pass
one detector and exiting bees another. The RFID
systems can provide data on bee life expectancy,
foraging time, and other crucial information
concerning hive dynamics. Schneider et al. (2012)
used the RFID technology to investigate sublethal
pesticide effects on bee colonies by exposing
workers from a small colony of about 2,000 bees
to contaminated sugar syrup at a feeder (>29,000
trips were observed but the number of bees tagged
was not reported). Detectors were set up at the hive
and the syrup feeder, and the effects of pesticide
exposure were measured as the return rate of
foragers from the feeder.
RFID detector and tag systems can be somewhat
expensive, and some care must be taken that neither
the tags nor the glue to attach them affects bee
behavior or survivorship. Tags are generally used
once, which adds to the cost of each replicate hive
and each experiment. Most RFID detectors require
modification of the hive entrance. The detectors
induce an electric current in the tags and need the tag
to pass at a short distance (often 5 mm or less); they
are thus sensitive to the orientation of the bee as it
crosses the detector threshold. Schneider et al. (2012)
modified the entrance to ensure proper orientation.
Because forager traffic is closely linked to colony
food intake and to pollination, it is a particularly
useful variable for researchers, beekeepers, and
growers. Burrill and Dietz (1981) found that while
changes in flight activity were directly proportional
to environmental temperature changes across the
observed temperature range, which was true for
changes in solar radiation only to a certain threshold
(0.66 langleys), above which flight activity de-
creased with increasing solar radiation. Danka and
18 W.G. Meikle and N. Holst
Beaman (2007) found no significant difference in
forager activity between the two bee races after
controlling for colony size, but did find significant
ambient temperature effects. Schneider et al. (2012)
found no effect on forager return success rate at
field-level doses of neonicotinoid pesticides, but
they did observe effects at higher doses.
2.5. Wireless networks
Sensors are often used in environmental and
agricultural monitoring, and data collection in
those fields is often managed using wireless
networks (Aqeel-ur-Rehman et al. 2014; Hart and
Martinez 2006; Ruiz-Garcia et al. 2009; Zerger et
al. 2010). Advances in technology and the increas-
ing availability of wireless network access even in
comparatively remote locations have made the use
of such networks a commonplace among re-
searchers, farmers, and land and wildlife managers.
Where wireless phone networks are not available,
transmitters are available to access satellite phone
networks. Aqeel-ur-Rehman et al. (2014) compare
different kinds of networks in terms of, among
other aspects, frequency band, cost, energy con-
sumption, and security. Zerger et al. (2010)
provided examples of how sensor networks have
been employed in three main application domains:
vegetation, animals, and soils.
Clearly, much potential exists for such networks
in apiculture. An excellent application of wireless
technology to create a network of beehives on scales
can be found on the internet at http://www.
bienenkunde.rlp.de. Data on hive weight and weight
change, as well as ambient temperature and r.h. are
available for over 70 sites across Germany, and those
data are updated every 5 min. Such region-wide
networks will likely prove very valuable for evalu-
ating the impact of regional effects, such as climate
or large-scale agricultural practices, which would be
difficult or impossible on a smaller scale.
3. DISCUSSION
Methods and technology for continuous
monitoring of beehives have changed since
Gates (1914), Milner (1921), and Hambleton
(1925), although their dedication to the task was
impressive. Modern studies involve more elec-
tronics with a higher sampling frequency for
more hive parameters. The use of sensors also
permits hive observation without disturbance;
gathering field data on colony growth and
phenology from frequent, invasive hive inspec-
tions, for example, to assess treatment effects,
can provoke bee defensive behavior (Breed et
al. 2004), enable robbing by other colonies
(Gary 1992), and lose, injure, or kill the queen
in addition to disrupting the hive environment.
Small, autonomous sensors, particularly those
linked to wireless networks, can provide much
information in real time with no disturbance.
Continuous monitoring has been applied to study
colony growth, activity, metabolic processes, genet-
ics, and behavior. One objective in continuously
monitoring has been to examine temporal patterns,
but many studies have tended more toward “proof of
concept” with the general application of the results
being hampered by a low number, or even lack, of
replicate bee colonies. Low replicate numbers can
be attributed largely to equipment costs, which are
not negligible; environmental chambers, respirome-
ters, electronic balances, accelerometers, RFID tags,
and associated equipment are expensive by most
standards (temperature sensors less so). Common
access to cheap and open-source electronics is quite
recent (Faludi 2010), and researchers have often
resorted to replication through time, if only for one
or a few hives, to compensate for the singularity of
their monitoring setup. As costs for electronics
decrease, the costs of replication will decrease and
thus become more widely applied.
Continuous monitoring involves, by definition,
conducting observations over time. While we
restricted the scope of this paper to studies with
two or more days of data collected at least hourly,
the length of time a variable could be measured in
a practical sense was in many ways determined by
the technology available and by the variable itself.
Continuous monitoring of any kind is limited
when it depends on constant human attention.
Gates (1914) and Hambleton (1925) exerted what
was surely considerable effort to monitor hives
hourly without interruption for just a few days at a
time, while Meikle et al. (2006, 2008) produced
hourly weight data for 16 months simply using
 Application of continuous monitoring of honeybee colonies
electronic data loggers. The quantity of data
produced increases rapidly with sampling frequen-
cy and the number of sensors and colonies
involved. Managing, and extracting useful infor-
mation from, continuous data from experiments
using large numbers of hives over extended
periods of time can be challenging. Analyses can
be simplified by exploiting patterns. Some vari-
ables, such as weight, temperature, and humidity,
have been found to have strong sinusoidal
components (Human et al. 2006; Meikle et al.
2008) owing to circadian rhythmicity; statistical
analyses can be conducted on curve parameters.
Monitoring bee colonies in the field can have
significant logistical requirements. In some cases,
technology has changed a great deal. To measure
temperature, Gates (1914) and Seeley (1974) used
mercury thermometers; Philips and Demuth
(1914), Milner (1921), and Owens (1971) used
electric thermocouples; whereas more recent re-
search has been conducted with small, inexpen-
sive, accurate, battery-powered electronic sensors
(e.g., Human et al. 2006). With battery power,
some sensors can last for months before memory
restrictions require a download and have sufficient
power to last for years. In other cases the changes
in technology have been less drastic. Electronic
scales sufficiently robust and precise for fieldwork
still tend to be expensive and heavy. In addition,
some sensors might not be readily transferred
among hives; as noted above, Bencsik et al. (2011)
embedded their accelerometers in hive walls, and
moving those accelerometers among hives might
be difficult. Detectors for forager traffic usually
require some modification of the hive entrance,
which may complicate rapid installation and
removal and may restrict bee activity.
Almost all of the sensors described here require
electricity. Commercially available sensors are
usually powered by (1) small watch-type batteries,
such as those sensors used by Human et al. (2006);
(2) wall current, which requires close proximity of
research hives to electric outlets; or (3) high amp-
hour batteries, often combined with an independent
power source, such as solar panels (e.g., Meikle et
al. 2006, 2008). Sensors such as load cells (with
indicators or signal conditioners) and RFID detec-
tors have higher requirements and are likely to need
19
12 or 24 V DC power or even AC power, which can
be provided by either wall current or batteries. How
long a given battery system will last before the
batteries need recharging depends on the number
and types of equipment drawing current from it,
including data loggers, wireless transmitters, and
transformers. Designing the system to only draw
power when a reading is taken would save energy,
but constantly rebooting the indicators and signal
conditioners should probably be avoided.
Researchers with remote hives must either change
batteries regularly or generate power using solar
panels or wind turbines. Solar power systems can be
placed, theoretically, almost anywhere, but do put
restrictions on the amount and form of power
available. The storage capacity and power draw on
the system will determine the size of the batteries
and energy generator required, taking into account
the need for some “autonomy” in the event that the
generator cannot function, such as cloudy weather
in the case of solar panels or calm in the case of
wind turbines. High-capacity, deep-cycle batteries
tend to be heavy and expensive.
4. CONCLUSIONS
Continuous monitoring will very likely become
a more common tool as both for research and in
practical apiculture as electronic components be-
come easier to deploy in the field, owing to small,
accurate, and robustly designed sensors. Data on
frequently measured variables, such as weight and
temperature, will likely be more thoroughly
exploited for information. Monitoring hive weight
or forager traffic prior to crop pollination would
allow beekeepers to observe hive health, and during
pollination, those data could provide a record of
quality control. Monitoring multiple variables
offers the possibility of synergy, as the information
that one method validates or augments information
gained simultaneously via another method.
Combining methods may also provide a way of
exploiting data from variables such as vibration, for
which the biological interpretation is not always
evident. Sound cues have already been the focus of
commercially available diagnostic tools for bee-
hives. Other anticipated improvements include the
following:
20 W.G. Meikle and N. Holst
1. Increased sampling frequency, for example,
may show changes over minutes or seconds
and thus explore colony behavior at differ-
ent time scales.
2. RFID tags are likely to be increasingly used
in bee research; these tags have commercial
application, such as for controlling inven-
tory, so that technology is likely to improve
substantially in the near future.
3. Cameras that detect ultraviolet and/or infrared,
as well as visible light are likely to be more
widely used as image analysis software im-
proves.
4. Bees have been found to exploit changes in
electric fields as a means of communication
(Greggers et al. 2013), opening up new
avenues for research and sensor development.
5. Sensors designed to detect the presence or
concentration of particular compounds may
help in disease or contamination monitoring.
ACKNOWLEDGMENTS
The authors would like to warmly thank A. Barron,
M. Bencsik, F. Linton, R. Mathew, A. Stilwell, M.
Weiss, and two anonymous reviewers for their insight-
ful comments and very helpful suggestions.
OPEN ACCESS
This article is distributed under the terms of the
Creative Commons Attribution License which per-
mits any use, distribution, and reproduction in any
medium, provided the original author(s) and the
source are credited.
Mise en place d’une surveillance en continu de
colonies d’abeilles
Poids de la ruche / température de la colonie /
humidité / activité des butineuses / vibration
Einsatzgebiete für eine kontinuierliche
Überwachung von Honigbienenvölkern
kontinuierliches Bienenvolkgewicht /
Bienenvolktemperatur / relative Feuchte /
Sammelaktivität / Vibration
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